ARTICLE IN PRESS

Quaternary International 169–170 (2007) 17–23

The proboscideans (Mammalia) from Mesoamerica

Joaquı́n Arroyo-Cabralesa,, Oscar J. Polacoa, César Lauritob, Eileen Johnsonc,
Marı́a Teresa Alberdid, Ana Lucı́a Valerio Zamorae
a
Laboratorio de Arqueozoologı´a ‘‘M. en C. Ticul Álvarez Solórzano’’, Subdirección de Laboratorios y Apoyo Académico, INAH,
Moneda # 16, Col. Centro, 06060 Mexico, D.F., Mexico
b
Instituto Nacional de Aprendizaje, Apartado Postal 203-2200, Coronado, Costa Rica
c
Museum of Texas Tech University, Lubbock, TX 79409-3191, USA
d
Museo Nacional de Ciencias Naturales, CSIC, José Gutiérrez Abascal 2, 28006 Madrid, Spain
e
Departamento de Historia Natural, Museo Nacional de Costa Rica, Apartado Postal 749-1000, San José, Costa Rica
Available online 17 January 2007

Abstract

Mesoamerica is the geographic region between northern México and southern Panamá. This region shows specific cultural and
biological features, both presently and in the past. The proboscideans (Mammalia, Proboscidea) that inhabited the region in a time range
from middle Miocene to late Pleistocene are outlined. Three families (Elephantidae, Gomphotheriidae, and Mammutidae) and six genera
currently are known from the region. The systematic position of the genus Stegomastodon is unresolved, whether monophyletic or
paraphyletic, and it may include two genera, Stegomastodon sensu stricto and Haplomastodon. The other five genera recognized for the
region are Gomphotherium (1 species), Rhynchotherium (3 species), Cuvieronius (3 species), Mammut (1 species), and Mammuthus
(2 species). The Great American Biotic Interchange played an important role for the migration of the proboscideans into South America.
The association between proboscideans and humans in Mesoamerica has been explored, but there is only evidence of an association with
mammoths exists.
r 2007 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction and regional setting
Mesoamerica is a culturally defined term in regards to
human culture and populations that inhabited the region
from northern México south to Panamá. This definition
has been expanded recently in Conservation Biology to
designate the biodiversity in the region limited to the north
by the Mexican–USA border and to the south by the
Panamanian–Colombian border. While those boundaries
are political, topographic and climatic conditions merge
this large area into a unique biodiversity unit. The northern
part of the unit is represented by México, while Central
America constitutes the southern part of the region.
This unit also may be reflected in the past, although large
climatic differences existed from the north (glaciations) to
the south (plateau rising ¼ faunal interchange). In this
region, a rich proboscidean fauna (Mammalia, Probosci-
Corresponding author. Tel.: +52 55 5522 4162; fax: +52 55 5522 3515.
E-mail address: arromatu5@yahoo.com.mx (J. Arroyo-Cabrales).
dea) occurred from Miocene to late Pleistocene times.
Three families inhabited México and northern Central
America, while the Gomphotheriidae were in the south and
migrated into South America during the Pleistocene
‘‘Great American Biotic Interchange’’ (Webb, 1978; Stehli
and Webb, 1985). Although Elephantidae is the only
surviving family at present, it was extirpated from the
Americas at the Pleistocene/Holocene boundary. Mammu-
tidae and Gomphotheriidae were extinct at the end of the
Pleistocene.
2. Discussion
The family Elephantidae is represented in Mesoamerica
by one genus with two species. The poorly known
Mammuthus hayi/meridionalis complex occurs during the
Middle Pleistocene with three putative localities in north-
ern and central México. Those identifications, however, are
questionable. Remains of Columbian mammoth (Elephan-
tidae, Mammuthus columbi) are the most widespread

1040-6182/$ - see front matter r 2007 Elsevier Ltd and INQUA. All rights reserved.
doi:10.1016/j.quaint.2006.12.017
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Quaternary fossils in the northern part of Mesoamerica
(Arroyo-Cabrales et al., 2003a). Several reasons may account
for that abundance, including sizeable mammoth popula-
tions at the end of the Pleistocene. A more practical reason is
that their enormous bones allowed people to recognize their
presence easily in the field. This situation, however, may
indicate the relative high abundance of this proboscidean in
the region. Mammoth remains also have been sought out
because of a possible association of those remains with
human activity. Lastly, a large volume of Quaternary
sediments is found in some regions, including the Basin of
México (Lorenzo and Mirambell, 1986; Arroyo-Cabrales et
al., 2003a, 2006). Although Pichardo (2005) has proposed the
erection of two subspecies for M. columbi, that scheme is not
supported by biological standards.
The Columbian mammoth has been recorded in México
since the 16th century and is known from most Mexican
states, with the exception of the Yucatan Peninsula (Fig. 1).
The largest concentration of localities is in the Basin of
México, and along the Mexican Plateau. Overall, 271
localities are known for the country (Arroyo-Cabrales
et al., 2003a). With such an extensive distribution, the usual
association of western camel (Camelops hesternus), ancient
bison (Bison spp.), horses (Equus spp.), and large predators
such as the Pleistocene lion (Panthera atrox) and short-
faced bear (Arctodus spp.) characterize a mammoth fauna.
To the south, a few records come from Chiapas (México)
(Arroyo-Cabrales et al., 2003b), Guatemala (Polaco,
unpublished data), Honduras (Webb and Perrigo, 1984),
Nicaragua (Espinoza, 1976), and Costa Rica (Laurito,
Fig. 1. Known distribution of Mammuthus (Proboscidea, Elephantidae) in Mesoamerica, with Mexican states and Central American countries
highlighted; note that localities are shown for state occurrence ranges in México, and specific localities in Central America, as available in the literature.
1988; Lucas et al., 1997). An isolated specimen from South
America (Guyana) reported in the 1930s has been
questioned and the specimen now is unavailable (Lucas
et al., 1997). At some Central American localities, the
associated fauna varies and tropical elements are incorpo-
rated, such as various ground sloths (Megalonyx,
Eremotherium), and toxodon (Mixotoxodon).
Most of the known Mesoamerican mammoth records
have been assigned to the late Pleistocene based on faunal
composition rather than radiocarbon dates. Those known
dates, either obtained directly from bone or from
associated sediments, range between 10,500 and 32,000 yr
BP (uncalibrated radiocarbon dates) (Arroyo-Cabrales
et al., 2003a; Gonzalez et al., 2003).
The American mastodon Mammut americanum repre-
sents the Mammutidae. This taxon is much less abundant
in the southern part of the region, with only one locality in
Honduras (Lucas and Alvarado, 1991) (Fig. 2). This form
is a more specialized proboscidean, feeding on herbs and
shrubs in temperate forests.
Sixteen mastodon localities are known from Mesoamer-
ica, 15 of those from México (Fig. 2). In three Mexican
localities, remains of the American mastodon are in
association with Columbian mammoth, and two are
possibly associated with human activity (Polaco et al.,
2001). An unexpected pattern emerges when the known
Mexican localities of American mastodon are plotted with
those of Pleistocene gomphotheres. Even with the possible
dating bias, these animals apparently are ecologically
parapatric.
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Fig. 2. Known distribution of Mammut (Proboscidea, Mammutidae) in Mesoamerica, with Mexican states and Central American countries highlighted;
note that specific localities are shown, as available in the literature.
Mammuthus and Mammut did not cross the Panamanian
land bridge, despite of the Guyanan record. One hypoth-
esis proposes that these two genera were highly specialized
feeders with habitat preferences not represented in the
Panamanian land bridge (Prado et al., 2005). The
contemporaneity of humans and proboscideans has been
examined only for mammoth (Arroyo-Cabrales et al.,
2006).
Gomphotheres (Gomphotheriidae) were the third major
proboscidean group that inhabited Mesoamerica. This
most successful, worldwide group appeared in North
America in the middle Miocene from Asian immigrants,
and lasted until the late Pleistocene (Lambert, 1996). As
this group crossed into South America during the Great
American Biotic Interchange event, it apparently did so
during the more arid glacial phase, when savanna habitats
extended broadly through tropical latitudes (Prado et al.,
2005). This family had the widest distribution for
proboscideans in the Americas. Recently, Alberdi and
Corona-M. (2005) reviewed most of the Gomphotheriidae
remains found in Mexican localities identified to genus
(at least 30 localities), focusing on their geographic
distribution. Several localities recorded the co-occurrence
of Rhynchotherium, Cuvieronius, and Stegomastodon.
Stegomastodon data primarily were from the deposits in
Valsequillo (State of Puebla).
Four genera occurred in northern Mesoamerica: two
longirostrine forms, Gomphotherium and Rhynchotherium;
and two brevirostrine ones, Stegomastodon and Cuvieronius
(Miller and Carranza-Castañeda, 2001; Polaco, 2003;
Alberdi and Corona-M., 2005; Corona-M. and Alberdi,
2006). Gomphotherium was the first of the gomphotheriids
19
to reach North America in the middle Miocene. It also was
recorded from southern México during the middle Miocene
(Barstovian) (Ferrusquı́a-Villafranca, 1990) and late Mio-
cene to early Pliocene in central México (Hemphillian)
(Miller and Carranza-Castañeda, 2001) (Fig. 3). Rhynch-
otherium was known only from the Americas, and probably
represents an indigenous radiation from the Gomphother-
ium stock (Lambert, 1996). Three species probably existed
in the Mesoamerican region (R. blicki, R. falconeri, and
R. tlascalae), but a taxonomic review is warranted.
Most of the records of Rhynchotherium are known only
from isolated individuals. A substantial population, how-
ever, comes from Honduras (Webb and Perrigo, 1984;
Miller and Carranza-Castañeda, 2001) and isolated dis-
coveries in Guatemala (Lucas and Alvarado, 1995) and
Costa Rica (Laurito and Valerio, 2007) (Fig. 3). This genus
may have been restricted to woodland savannas (Lambert,
1996).
Stegomastodon and Cuvieronius are known from both
North and South America. Stegomastodon ranges from
early Blancan to early Irvingtonian. Although the genus
was considered as the more specialized grazer within the
American gomphotheres, it has been redefined as a mixed
feeder with tendencies toward both browsing and grazing
(Prado et al., 2005). This feeding habit indicates that the
genus may have been adapted to open grasslands with
warm to temperate conditions.
Cuvieronius first appeared in the North American
Blancan, and persisted until late Pleistocene. The known
geographic distribution of the genus in México comprised
mainly the central and southern states, with a few records
from northern México (Montellano-Ballesteros, 2002). It
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Fig. 3. Known distribution of Gomphotherium and Rhynchotherium (Proboscidea, Gomphotheriidae) in Mesoamerica, with Mexican states and Central
American countries highlighted; note that specific localities are shown, as available in the literature for Gomphotherium, while shaded states in México are
provided for Rhynchotherium.
also was known for most of Central America, from
Guatemala to Costa Rica (Lucas et al., 1997) (Fig. 4).
The genus was a mixed feeder, inhabiting high grasslands
with cold to temperate climatic conditions (Prado et al.,
2005). As many as three species may be recognized in the
region: C. hyodon, C. oligobunis, and C. tropicus (Laurito,
1988; Montellano-Ballesteros, 2002; Prado et al., 2005).
The Mesoamerican brevirostrine genera have had a very
complex taxonomic history that has comprised between
two and three genera. When considered, the third genus is
Haplomastodon. The presence of this form may be exclusive
to South America. Its postulated presence, however, may
mean that the Central America form of Stegomastodon
evolved into Haplomastodon in South America. Two
species are recognized for Stegomastodon in South Amer-
ica, Stegomastodon platensis and Stegomastodon waringi
(Prado et al., 2005), while for North America, Stegomas-
todon primitivus and Stegomastodon mirificus are noted
(Osborn, 1929). Another species of Stegomastodon, how-
ever, may be found in the late Pleistocene of central México
(Alberdi et al., 2002). A clear need exists for a detailed
taxonomic review of the brevirostrine gomphotheres
associated with Stegomastodon sensu lato.
Prado et al. (2005) are in agreement with Simpson and
Paula Couto (1957) that the genus Haplomastodon cannot
be differentiated clearly from Stegomastodon. They believe
that Haplomastodon (Hoffstetter, 1950), is a junior
synonym of Stegomastodon (Pohlig, 1912). Ficcarelli
et al. (1997), however, consider the genus Haplomastodon
to be present in the tropical zone of South America, and
Stegomastodon in the South American prairies, from Brazil
throughout the Pampean Region.
This issue has not been resolved, and two of the co-
authors of this paper have contradictory views. Alberdi
regards the remains from Haplomastodon and Stegomasto-
don as very similar. This similarity is a taxonomic problem
that requires an in-depth study of similarities and
differences, as well as recovering the ancestral form to
define their relationships and any synonymies. Haplomas-
todon is not found in Mesoamerica or North America. It
was first proposed at the subgeneric level on variable
characteristics (Hoffstetter, 1950). Hoffstetter (1952) raised
it to the genus level without a detailed explanation of the
diagnostic generic characters. In order to validate any
hypothesis, all of the remains assigned to Stegomastodon in
both North and South America would need to be studied,
and then a decision made as to the number of taxa
involved. If two genera were found, then Haplomastodon
would be recognized.
On the other hand, Laurito points out the differences in
distribution. Cuvieronius has a wide, continuous distribu-
tion from North America, Mesoamerica, and northern
South America. Stegomastodon distribution, however, is
discontinuous, with a large gap in southern México and
Central America. The discovery of several proboscideans in
the region during the 19th and 20th centuries (Leidy, 1859;
Olson and McGrew, 1941; Webb and Perrigo, 1984;
Laurito, 1988; Lucas and Alvarado, 1991; Cisneros, 2005;
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Fig. 4. Known distribution of Cuvieronius (Proboscidea, Gomphotheriidae) in Mesoamerica, with Mexican states and Central American countries
highlighted; note that specific localities are shown, as available in the literature.
Pearson, 2005; Laurito and Valerio, 2007) could be the
basis for a study of the paraphyletic relationships within
the American brevirostrine gomphotheres. As for the
origin of the South American Stegomastodon, an early
gomphothere arrival may have occurred on the southern
continent, as either a species within Rhynchotherium or
Cuvieronius, which eventually evolved into Stegomastodon.
Cozzuol (personal communication, 2005) proposes Ama-
huacatherium peruvium as the possible ancestor for the
group. It is a late Miocene tetrabelodont mastodont from
Perú whose dentition is morphologically similar to that of
Rhynchotherium and the South American stegomastodonts
(Campbell et al., 2000). The minimal age for the
Amazonian locality is around 9.0170.28 million years
(Campbell et al., 2001). Such an isolated date for
supporting the geologic history of the wide Amazonian
Basin, however, remains to be confirmed (Alberdi et al.,
2004).
A systematic study of the North and South American
remains identified as Stegomastodon is warranted based on
dental and cranial characters to assess the phylogeny and
decide whether the genus Stegomastodon was monophyletic
or paraphyletic. A distribution map is not provided for the
genus because of the taxonomic uncertainty. A human
association with American mastodon and with gom-
photheres has not been confirmed for Mesoamerica.
21
Overall, are tentative records exist for at least nine
species (possible one more), six genera, and three families
within the order Proboscidea for Mesoamerica. The known
species occurred from the Middle Miocene to Late
Pleistocene, with the largest abundance during the Pliocene
(Fig. 5).
3. Concluding remarks
Four proboscidean genera may have been contempora-
neous at the end of the Pleistocene in northern Mesoamer-
ica. Furthermore, at some localities, mammoth and
American mastodon have been found in the same deposits,
or mammoth and gomphothere, but not American
mastodon and gomphothere together. This situation
reflects the apparent parapatric relationship.
Detailed taxonomic and systematic studies are required
to define clearly the number of species for mammoths or
genera for gomphotheres that occurred during late Tertiary
and early Quaternary in the Americas. A review of all of
the specimens held in museums is warranted to allow a
complete understanding of the Mesoamerican probosci-
deans. Stratigraphic and radiometric-controlled excava-
tions are required to enhance the proboscidean records for
Mesoamerica as well.
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22 J. Arroyo-Cabrales et al. / Quaternary International 169–170 (2007) 17–23
Fig. 5. Geologic range chart for the proboscidean species known for Mesoamerica. Those for which there are more uncertainties about their time
occurrence are marked with a question sign (?).
Finally, at present, the proboscidea information for
Mesoamerica is incomplete, but the records show great
potential for increasing and enhancing such knowledge.
Furthermore, their study could help in understanding the
evolutionary patterns and extinction causes for the unique
order Proboscidea in the New World.
Acknowledgements
The authors thank Mario Cossuol (Universidad de
Rondonia, Brazil) for sharing unpublished information.
Felisa Aguilar kindly prepared the maps. Several institu-
tions supported parts of the research: Comisión Nacional
para el Conocimiento y Uso de la Biodiversidad (CON-
ABIO), Consejo Nacional de Ciencia y Tecnologı́a (CON-
ACYT), the Subdirección de Laboratorios y Apoyo
Académico, Instituto Nacional de Antropologı́a e Historia,
México, and the Museum of Texas Tech University.
Johnson’s participation represents part of ongoing regional
research of the Lubbock Lake Landmark into Quaternary
grasslands and ecological changes in the Americas under
the auspices of the Museum of Texas Tech University.
References
Alberdi, M.T., Corona-M., E., 2005. Revisión de los gonfoterios en el
Cenozoico tardı́o de México. Revista Mexicana de Ciencias Geológi-
cas 22, 246–260.
Alberdi, M.T., Polaco, O.J., Juárez Woo, J., Arroyo-Cabrales, J., 2002.
Presencia de Stegomastodon (Mammalia: Gomphotheridae) en el
Pleistoceno final de Jalisco, México. Abstract, VIII Congreso
Mexicano de Paleontologı́a, 12 al 16 de noviembre del 2002, Museo
de Paleontologı́a ‘‘Ing. Federico Solórzano Barreto’’, Guadalajara,
Jalisco, 28 pp.
Alberdi, M.T., Prado, J.L., Salas, R., 2004. The Pliestocene Gomphother-
iidade (Proboscidea) from Peru. Neues Jarbuch fur Geologie und
Palaontologie Abhandlungen 231, 423–452.
Arroyo-Cabrales, J., Polaco, O.J., Johnson, E., Guzmán, A.F., 2003a. The
distribution of the genus Mammuthus in México. In: Reumer, J.W.F.,
De Vos, V., Mol, D. (Eds.), Advances in Mammoth Research
(Proceedings of the Second International Mammoth Conference,
Rotterdam, May 16–20, 1999), DEINSEA 9, pp. 27–39.
Arroyo-Cabrales, J., Polaco, O.J., Aguilar-Arellano, F.J., 2003b. Remains
of Mammuthus housed in the collections of the Instituto Nacional de
Antropologı́a e Historia, México. In: Reumer, J.W.F., De Vos, V.,
Mol, D. (Eds.), Advances in Mammoth Research (Proceedings of the
Second International Mammoth Conference, Rotterdam, May 16–20,
1999), DEINSEA 9, pp. 17–25.
Arroyo-Cabrales, J., Polaco, O.J., Johnson, E., 2006. A preliminary view
of the coexistence of mammoth and early peoples in México.
Quaternary International 142–143, 79–86.
Campbell, K.E., Frailey, C.D., Romero, L., 2000. The Late Miocene
Gomphothere Amahuacatherium Peruvium (Proboscidea: Gom-
photheriidae) from Amazonian Peru: implications for the great
American faunal interchange. INGEMMET, 152pp.
Campbell, K.E., Heizler, M., Frailey, C.D., Romero, L., Prothero, D.R.,
2001. Upper Cenozoic chronostratigraphy of the southwestern
Amazon Basin. Geology 29, 595–598.
Corona-M, E., Alberdi, M.T., 2006. Two new records of Gomphother-
iidae (Mammalia, Proboscidea) in Southern Mexico and some
biogeographic implications. Journal of Paleontology 80, 357–366.
Cisneros, J.C., 2005. New Pleistocene vertebrate fauna from El Salvador.
Revista Brasileira de Paleontologı́a 8, 239–255.
Espinoza, J., 1976. Evaluaciones arqueológicas en ‘‘El Bosque’’. Instituto
Geográfico Nacional, Managua, Ministerio de Obras Públicas 1,
pp. 22–45.
Ferrusquı́a-Villafranca, I., 1990. Biostratigraphy of the Mexican con-
tinental Miocene: part II, the southeastern (Oaxacan) faunas.
Paleontologı́a Mexicana, Instituto de Geologı́a, Universidad Nacional
Autónoma de México, vol. 56, pp. 57–109.
Ficcarelli, G., Azzaroli, A., Bertini, A., Coltorti, M., Mazza, P.,
Mezzabotta, C., Moreno Espinosa, M., Rook, L., Torre, D., 1997.
Hypothesis on the cause of extinction of the South American
Mastodonts. Journal of South American Earth Sciences 10, 29–38.
Gonzalez, S., Jiménez-López, J.C., Hedges, R., Huddart, D., Ohman, J.C.,
Turner, A., 2003. Earliest humans in the Americas: new evidence from
México. Journal of Human Evolution 44, 379–387.
Hoffstetter, R., 1950. Observaciones sobre los mastodontes de Sud
América y especialmente del Ecuador. Haplomastodon, subgen. nov. de
Stegomastodon. Publicaciones Escuela Politécnica Nacional 1, 1–49.
 ARTICLE IN PRESS
J. Arroyo-Cabrales et al. / Quaternary International 169–170 (2007) 17–23
Hoffstetter, R., 1952. Les mammifères pleistocenes de la republique de
l’equateur. Mémoires Société Géologique de France 66, 1–391.
Lambert, W.D., 1996. The biogeography of the gomphotheriid probosci-
deans of North America. In: Shoshani, J., Tassy, P. (Eds.), The
Proboscidea. Evolution and Palaeoecology of Elephants and Their
Relatives. Oxford University Press, Oxford, pp. 143–148.
Laurito, C.A., 1988. Los proboscideos fósiles de Costa Rica y su contexto
en la América Central. Vı́nculos, Revista de Antropologı́a del Museo
Nacional de Costa Rica 14, 29–58.
Laurito, C.A., Valerio, A.L., 2007. First record of Rhynchotherium blicki
(Frick, 1933) for the late Cenozoic of Costa Rica. Revista Geológica de
América Central 33, 75–82.
Leidy, J., 1859. On mastodon tooth from Honduras. Proceedings of the
Academy of Natural Sciences of Philadelphia 2, 91.
Lorenzo, J.L., Mirambell, L., 1986. Mamutes excavados en la Cuenca de
México (1952–1980), Cuaderno de Trabajo, Departamento de
Prehistoria, Instituto Nacional de Antropologı́a e Historia, México
32, 1–151.
Lucas, S.G., Alvarado, G.E., 1991. El hallazgo más austral de un Mammut
americanum: el caso del mastodonte de San Pedro Sula, Honduras.
Revista Geológica de América Central 13, 85–89.
Lucas, S.G., Alvarado, G.E., 1995. El proboscideo Rhynchotherium blicki
(Mioceno Tardı́o) del oriente de Guatemala. Revista Geológica de
América Central 18, 19–24.
Lucas, S.G., Alvarado, G.E., Vega, E., 1997. The Pleistocene mammals of
Costa Rica. Journal of Vertebrate Paleontology 17, 413–427.
Miller, W.E., Carranza-Castañeda, O., 2001. Late Cenozoic mammals
from the basins of central México. Bollettino della Società Paleonto-
logica Italiana 40, 235–242.
Montellano-Ballesteros, M., 2002. New Cuvieronius finds from the
Pleistocene of Central México. Journal of Paleontology 76, 578–583.
23
Olson, E.C., McGrew, P.O., 1941. Mammalian fauna from the Pliocene of
Honduras. Bulletin of the Geological Society of America 52,
1219–1244.
Osborn, H.F., 1929. New Eurasiatic and American proboscideans.
American Museum Novitates 393, 1–23.
Pearson, G.A., 2005. Late Pleistocene Megafaunal deposits on the Isthmus
of Panama and their Paleoenvironmental implications. Caribbean
Journal of Science 41, 1–13.
Pichardo, M., 2005. Taxonomic revision of central Mexican mammoths in
Paleoindian sites. Anthropologische Anzegier 63, 409–413.
Polaco, O.J., 2003. Los proboscidios de México. In: González, G.A.H., De
Stéfano, F.A. (Eds.), Fósiles de México. Coahuila, una ventana a
través del tiempo. Gobierno del Estado de Coahuila, México, p. 179.
Polaco, O.J., Arroyo-Cabrales, J., Corona-M., E., Oliva-López, J.G.,
2001. The American Mastodon Mammut americanum in México. In:
Cavarretta, G., Gioia, P., Mussi, M., Palombo, M.R. (Eds.), La Terra
degli Elefanti. Atti del 11 Congresso Internazionale. Roma, 16–20
Ottobre 2001. Consiglio Nazionale delle Ricerche, Roma, Italia,
pp. 237–242.
Prado, J.L., Alberdi, M.T., Azanza, B., Sánchez, B., Frassinetti, D., 2005.
The Pleistocene Gomphotheriidae (Proboscidea) from South America.
Quaternary International 126/128, 21–30.
Simpson, G.G., Paula Couto, C., 1957. The Mastodonts of Brazil. Bulletin
of the American Museum of Natural History 112, 125–190.
Stehli, F.G., Webb, S.D. (Eds.), 1985. The Great American Interchange.
Plenum Press, New York.
Webb, S.D., 1978. A history of savanna vertebrates in the new World. Part
II: South America and the great interchange. Annual Review of
Ecology and Systematics 9, 393–426.
Webb, S.D., Perrigo, S.C., 1984. Late cenozoic vertebrates from honduras
and El Salvador. Journal of Vertebrate Paleontology 4, 237–254.